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1 Colin Pittendrigh in his final years at Princeton University The manuscript associated with these remarks is an abbreviated version of my Ph.D. dissertation, earned in 1969 under the guidance of Colin Pittendrigh (Pitt) at Princeton University. The manuscript was not published, or more accurately was rejected, by Science magazine in 1976. I chose the subject because I was interested in the ecology of flies of the genus Drosophila, and I wanted to generate evidence that natural environmental conditions are a source of selection for the well known circadian eclosion rhythm of these animals. All species of Drosophila tested have a circadian eclosion rhythm that causes adult flies to emerge from the pupal case only in the first few hours after dawn. I selected one species of Drosophila native to the western U.S., and a second species native to the eastern U.S, and exposed both to natural and extreme saturation deficit regimes. Saturation deficit is a measure of the evaporative power of the atmosphere at a particular temperature and relative humidity. I wanted to find out if Drosophila in nature more successfully eclose in the early morning because that is the daylight time when the saturation deficit is usually at a minimum. I found this to be true for both species when exposed to their natural environment. In the laboratory, I found a positive relationship between eclosion success and low saturation deficits. Pitt steadily encouraged me in this work, reviewed the manuscript and made helpful suggestions before I submitted it for publication. I arrived at Princeton in the fall of 1965, eager to meet my new scientific mentor, Colin Pittendrigh. I had developed an interest in circadian rhythms as an undergraduate, and decided he was the person I wished to work with. I closely watched and learned from him, the most important person in my professional development. As it happened, Pitt had just become Dean of the Graduate School, a major administrative job with offices in historic Nassau Hall, far removed from his lab physically and psychologically. His lab was very busy, with several graduate students, post docs, and full- time laboratory assistants and technicians. The suite of rooms in 19 th Century Guyot Hall was a beehive of scientific activity, or perhaps I should say a fly swarm. Much of the research on circadian rhythms revolved around one organism, Drosophila pseudoobscura,a western North American species of fruit fly. Among the students in the lab at that time were the late Arthur Winfree of the University of Arizona, a mathematical biologist and MacArthur Fellow who wrote The Geometry of Biological Time, and Jerry Feldman of Swarthmore College, who has published many papers on molecular circadian clocks. There were often visitors to our laboratory. Some were colleagues from other universities. On occasion, there would be reporters and film crews interviewing Pitt on subjects as diverse as the possibility of extraterrestrial life, and federal support of scientific research. We were once visited by Jürgen Aschoff, who together with Pitt and Erwin Bünning laid the groundwork for modern studies of chronobiology. Aschoff met individually with every researcher in the lab, and listened with interest as we each explained our work. My first research idea was a hypothesis that the blind cave dwelling crustacean, Orchonectes pellucidus, long isolated underground in Mammoth Cave, Kentucky, would have retained circadian rhythms. I planned to bring animals into constant laboratory conditions, and measure their cyclical physical and physiological functions. I spent my first

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Page 1: Colin Pittendrigh in his final years at Princeton University · 2019. 8. 21. · Colin Pittendrigh in his final years at Princeton University The manuscript associated with these

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ColinPittendrighinhisfinalyearsatPrincetonUniversity

ThemanuscriptassociatedwiththeseremarksisanabbreviatedversionofmyPh.D.dissertation,earnedin1969undertheguidanceofColinPittendrigh(Pitt)atPrincetonUniversity.Themanuscriptwasnotpublished,ormoreaccuratelywasrejected,bySciencemagazinein1976.IchosethesubjectbecauseIwasinterestedintheecologyoffliesofthegenusDrosophila,andIwantedtogenerateevidencethatnaturalenvironmentalconditionsareasourceofselectionforthewellknowncircadianeclosionrhythmoftheseanimals.AllspeciesofDrosophilatestedhaveacircadianeclosionrhythmthatcausesadultfliestoemergefromthepupalcaseonlyinthefirstfewhoursafterdawn.IselectedonespeciesofDrosophilanativetothewesternU.S.,andasecondspeciesnativetotheeasternU.S,andexposedbothtonaturalandextremesaturationdeficitregimes.Saturationdeficitisameasureoftheevaporativepoweroftheatmosphereataparticulartemperatureandrelativehumidity.IwantedtofindoutifDrosophilainnaturemoresuccessfullyecloseintheearlymorningbecausethatisthedaylighttimewhenthesaturationdeficitisusuallyataminimum.Ifoundthistobetrueforbothspecieswhenexposedtotheirnaturalenvironment.Inthelaboratory,Ifoundapositiverelationshipbetweeneclosionsuccessandlowsaturationdeficits.Pittsteadilyencouragedmeinthiswork,reviewedthemanuscriptandmadehelpfulsuggestionsbeforeIsubmitteditforpublication.IarrivedatPrincetoninthefallof1965,eagertomeetmynewscientificmentor,ColinPittendrigh.Ihaddevelopedaninterestincircadianrhythmsasanundergraduate,anddecidedhewasthepersonIwishedtoworkwith.Icloselywatchedandlearnedfromhim,themostimportantpersoninmyprofessionaldevelopment.Asithappened,PitthadjustbecomeDeanoftheGraduateSchool,amajoradministrativejobwithofficesinhistoricNassauHall,farremovedfromhislabphysicallyandpsychologically.Hislabwasverybusy,withseveralgraduatestudents,postdocs,andfull-timelaboratoryassistantsandtechnicians.Thesuiteofroomsin19thCenturyGuyotHallwasabeehiveofscientificactivity,orperhapsIshouldsayaflyswarm.Muchoftheresearchoncircadianrhythmsrevolvedaroundoneorganism,Drosophilapseudoobscura,awesternNorthAmericanspeciesoffruitfly.AmongthestudentsinthelabatthattimewerethelateArthurWinfreeoftheUniversityofArizona,amathematicalbiologistandMacArthurFellowwhowroteTheGeometryofBiologicalTime,andJerryFeldmanofSwarthmoreCollege,whohaspublishedmanypapersonmolecularcircadianclocks.Therewereoftenvisitorstoourlaboratory.Somewerecolleaguesfromotheruniversities.Onoccasion,therewouldbereportersandfilmcrewsinterviewingPittonsubjectsasdiverseasthepossibilityofextraterrestriallife,andfederalsupportofscientificresearch.WewereoncevisitedbyJürgenAschoff,whotogetherwithPittandErwinBünninglaidthegroundworkformodernstudiesofchronobiology.Aschoffmetindividuallywitheveryresearcherinthelab,andlistenedwithinterestasweeachexplainedourwork.Myfirstresearchideawasahypothesisthattheblindcavedwellingcrustacean,Orchonectespellucidus,longisolatedundergroundinMammothCave,Kentucky,wouldhaveretainedcircadianrhythms.Iplannedtobringanimalsintoconstantlaboratoryconditions,andmeasuretheircyclicalphysicalandphysiologicalfunctions.Ispentmyfirst

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yearplanningthatproject,onlytodiscoverthatgraduatestudentThomasJeglahadjustcompletedthoseveryexperimentsforhisdissertationatYale.Hepublishedhisresults,showingthatthecrayfishdoexpressfreerunningcircadianrhythmsinconstantconditions,intheJournalofExperimentalZoologyin1968.Ihadtostartoverwithanotheridea.Pittasalwayswassupportive,andwebegantodiscussresearchalternatives.IformulatedaquestionthatwoulduseD.pseudoobscura.Iwantedtomakeupforlosttime,andthischoicemeantthatIcouldtakeadvantageofequipmentandproceduresalreadyinuseinhislaboratory.Iwasoffandrunning.Myexperimentaldesignrequiredexposingflypupaetonaturalenvironmentalconditions.IdidthisnearPrincetonusingasmalllaboratorybuiltformeinsideacenturyoldfarmhousebarnonpropertyownedbytheUniversity.Thisfieldlabmusthavebeenacostlyconstructionproject.IdonotknowbecauseafterIaskedaboutit,inshortorderitwasbuiltformeaspermyspecifications.ItwasPitt,ofcourse,whoarrangedandpaidforthat.Henevertoldmehow.ThefollowingyearIneededtotakemyexperimentalfliestoawesternU.S.climate,wheretheynaturallyoccur,tofindouthowpreciselythewellstudiedcircadianeclosionrhythmexhibitedinourPrincetonlaboratoryisentrainedbynaturaltemperatureandlightcycles.PittsentmetotheRockefellerUniversityinNewYorkCitytoconferwithTheodosiusDobzhansky,atthetimethegrandoldmanofevolutionarygenetics.PittpointedouttomethatDobzhanskywasmyintellectualgrandfather,sincehehadbeenPitt’smajorprofessoratColumbia.Pittreferredtohimas“theoldboy.”DobzhanskyhadbeenworkingformanydecadeswithDrosophila,andhaddonesointheAmericanWest.Thefirstthinghesaidtomewas“Sohereistheyoungmanwhodoesnotsignhisletters.”Ihadtypedmyletterstohim,andinmycallowyearsIdidnotrealizeitwasgoodformtoincludeasignature.Hemadethisremarkwithakindlysmileonhisface,becauseDr.Dobzhanskywasamarkedlycourteous,indeedcourtly,personwhogreetedmeasacolleague.HelistenedwithinterestasIexplainedmyresearch,andsoughthisadvice.InhispronouncedRussianaccent,headdressedmeas“MeesterQuinn.”Heshowedmearoundhislab,whereanumberofresearchers,whowerefamiliartomeonlyfromtheirpublications,explainedtheirresearchprograms.Dobzhanskytheninvitedmetopresentaseminaronmyresearchtohisassembledlaboratorycolleaguesandstaff.Iaskedhimwhen,expectingalaterdate,andhereplied“thisafternoonatfour,”twohourslater.HowcouldIsayno,eventhoughIwascompletelyunpreparedandquitenervousaboutanaudiencewhohadbeenfamouslystudyingDrosophilafordecades?Isomehowgatheredmywitsandmadeacoherentpresentation,witharmwavinginplaceofslides.AfterwardsDr.DobzhanskyinvitedmetodinnerwithhimandhispostdocsattheRockefellerUniversity,aplacewheremenwererequiredtodressfordinnerinacoatandtie.Icamethatdaydressedasahumblegraduatestudent.Thegooddoctorarrangedforoneofhispost-docs,atallman,toloanmethepropergarb,soIateandconversedwiththesleevesofmycoatjacketrolledup.Veryelegant.Dr.DobzhanskyrecommendedthatIdomyfieldworkattheSouthwesternResearchStation,afacilityinArizonaoperatedthenandnowbytheAmericanMuseumofNaturalHistory.Hehadworkedthere,wherehewasrememberedfortravellingaroundtohisfieldsitesonhorseback.TheBiologyDepartmentpaidformytravelexpenses,butnotforahorse,andPitthadtheexperimentalmachineryIneededbuiltandshippedtoArizona.

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ColinPittendrighhadoneofthemostremarkablescientificmindsthatIhaveeverencountered.Hegraspedcomplexideasimmediately,andoftenforesawtheirimplicationsforresearchjustasquickly.Ialwayscarefullypreparedformymeetingswithhiminhisdistantadministrativeoffices,becauseIknewthathecouldprocessandanalyzewhateverIsaidasquicklyasIcouldexplainmyself.Afterourmeetings,Iwouldsitdownandwriteouteverythinghehadsaid,whileitwasfreshinmymind.Hisglitteringverbalandanalyticalabilitieswereequallymatched.Whenhespokeformallyhewaselegant,andheknewhowtotailorhisremarkstohisaudience.Heenjoyedexplainingevolutioninthegeneralsense.IsawhimdothatatapubliclectureinthePrincetonPublicLibrary.Iwastakenbyhislecturebecausehestatedhisideassoclearlyforaverygeneralaudience.Onanotheroccasion,Isawhimexplainevolutionaryprinciples,inonehour,toaverylargeundergraduateclass.Hisremarksweresoengagingthatwhenheconcludedtheentireclassstoodandgavehimastandingovation.ThatwastheonlytimeIsawthathappeninanyPrincetonundergraduateclass.AsIwaspushingtowardthecompletionofmydissertationinthespringof1969,IlearnedthatPittwasleavingPrinceton,aswasI.HemovedtoStanfordUniversitywherehecontinuedhisresearch,andassumedanewsetofimportantadministrativeduties.Heofcoursecontinuedhisscientificresearch.Heoncetoldmethathewasconcernedthathewaslivingonhis“scientificcapital,”implyingthathisadministrativeresponsibilitieswerecausinghimtoneglectscientificresearch.IdidnotseethathappenduringhisfinalPrincetonyears.IfeelextremelyfortunatetohavestudiedunderPitt.Hewassupportive,andbyhisexamplehehelpedmelearntothinkandworkasascientist.RonaldD.QuinnPh.D.PrincetonUniversity,1969Dr.QuinnisProfessorEmeritusfromtheDepartmentofBiologicalSciencesatCalPolyPomona(akaCaliforniaStatePolytechnicUniversity,Pomona)

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